Social organization of the Alaotran gentle lemur (Hapalemur griseus alaotrensis)

Knowledge of the social organization of lemurs is still limited for most species. Where there is sufficient information, it has been shown that lemur social organization differs in essential points from that of other primates. In the field study reported here, demographic structure and life-history processes were investigated in order to characterize the social organization of the Alaotran gentle lemur (Hapalemur griseus alaotrensis). Data were obtained through captures and observations. Alaotran gentle lemurs were found in small groups of up to nine individuals. Although most groups contained just one breeding female, a substantial proportion of groups (35%) had two breeding females. Therefore, Alaotran gentle lemurs cannot be classed as being organized in monogamous family groups. An extended birth season was found, and groups with two breeding females had significantly higher breeding output per adult than groups with a single adult female. Limited data suggest that females emigrate from their natal group while still subadult, whereas males can stay in the natal group until they are fully grown and disperse as adults. Variability in group composition, significantly higher reproductive output per adult in groups with two breeding females, and delayed dispersal of males suggest that Alaotran gentle lemurs pursue a resource-defense mating strategy, rather than a female-defense mating strategy. The suggestion that extant social lemurs may have evolved from a monogamous system, could explain the differences between lemur social systems and those of other primates.     

Monogamy as a Rule Rather than Exception in Nocturnal Lemurs: the Case of the Fat-tailed Dwarf Lemur, Cheirogaleus medius

According to present hypotheses on the evolution of life history traits and social systems in Malagasy lemurs, nocturnality and infant parking are associated with a solitary lifestyle and a polygynous mating system. However, theoretically extreme seasonality of reproduction could limit the number of females that can be monopolized by a given male and thus hinder the evolution of polygyny. The aim of this ongoing study is to test these contrasting expectations by looking at the social and mating system of the fat-tailed dwarf lemur Cheirogaleus medius . This species hibernates for up to 7 mo, so that time for breeding and raising offspring is extremely limited.
A mark-recapture study in western Madagascar was combined with observations and radio-tracking of 36 individuals during the rainy seasons from 1995 to 1998. According to these data, fat-tailed dwarf lemurs live in permanent sleeping groups consisting of a male and a female (n = 8) or one male and two females (n = 1). One or two offspring from the previous year were frequently observed to sleep together with an adult pair. Members of each sleeping group were the exclusive users of their nest holes and home ranges. During the birth season, males and females took turns at baby-sitting their offspring. Females without paternal help were unable to raise their offspring successfully. Since females did not exhibit oestrus synchrony, the ultimate selective factor favouring pair-living could be obligate paternal investment. The results, together with the lack of sexual size dimorphism and relatively small testis size, suggest that the fat-tailed dwarf lemur lives in family groups with a monogamous mating system. A review of the mating systems of nocturnal lemurs shows that monogamy appears to be the rule rather the exception.     

Central males instead of multiple pairs in redfronted lemurs, Eulemur fulvus rufus (Primates, Lemuridae)?

In group-living Malagasy primates (Lemuriformes), certain demographic and morphological traits deviate both from theoretical expectations derived from sexual selection theory and from patterns in better-known anthropoid primates. Lemurs lack sexual dimorphism in body and canine size and live in relatively small multimale-multifemale groups with, on average, even adult sex ratios, despite a polygynous mating system. In addition, the majority of gregarious lemurs are cathemeral, that is, they are regularly active both day and night. The social system of cathemeral lemurs has been considered an adaptation to their activity pattern. If so, groups should consist of multiple pairs that aggregate during diurnal activity and range separately at night in response to predation and infanticide risk, respectively. Moreover, mating privileges should exist between pair partners. We tested this hypothesis with data collected on two groups of wild redfronted lemurs in western Madagascar during 1023 focal animal hours spanning 4 months including the mating season. In addition, data on spatial relations were collected on 12 nights. We found no differences in group cohesion between day and night and no behavioural evidence for multiple male-female pairs. Instead, one male in each group monopolized social interactions with all females. However, females copulated with all resident males, although most often with the central male. Thus, basic predictions of the cathemerality hypothesis are unsupported. Multiple matings and indications of oestrous synchrony can be viewed as female counterstrategies against infanticide, whereas reproductive strategies of redfronted lemur males remain obscure. Copyright 1999 The Association for the Study of Animal Behaviour.     

Group encounters and territoriality in wild Alaotran gentle lemurs (Hapalemur griseus alaotrensis)

During a 3 month field study, 18 group encounters between four groups of Alaotran gentle lemurs (Hapalemur griseus alaotrensis) were observed in the Lake Alaotra marshland in Madagascar. Behaviors observed during group encounters are described, and quantitative data on intergroup interactions and ranging patterns are discussed in the context of territoriality. Intergroup interactions varied in their intensity, ranging from visual monitoring, scent marking, and display locomotion to penetrating the neighboring range. A quarter (27%) of the encounters were aggressive, involving chases and confrontation displays, and no affiliative interactions were observed. Both adult males and females were involved in intergroup encounters, with males playing more active roles. All encounters took place in the small overlapping areas of neighboring home ranges. The outcome of an aggressive encounter was determined by its location, with the resident group always driving out the intruding one. The Alaotran gentle lemur actively advertised and consistently defended a large area of its home range against intrusion of neighboring groups and can therefore be considered territorial. Am. J. Primatol. 46:251–258, 1998. © 1998 Wiley-Liss, Inc.     

Agonistic and grooming behavior of captive crowned lemurs (Lemur coronatus) during the breeding season

Captive breeding of rare and endangered prosimians may be enhanced by an increased understanding of all aspects of their seasonal mating behavior. In this study, the agonistic and grooming behaior of captiveL. coronatus were studied during their annual breeding season. Between October and March, selected aspects of agonistic and grooming behavior of four male-female groups were recorded. Males initiated significantly more allogrooming than females. The frequency of this behavior pattern increased significantly in the weeks of estrus. Females performed significantly more agonistic behavior patterns than males. The mean frequency of agonistic interactions decreased significantly during the breeding season. In successfully reproducing groups males showed much less agonism towards females, than they did in unsuccessful groups. Inter-specific comparisons demonstrated that the patterns of agonistic interactions during the breeding season are very different among closely related species of lemurids. The conclusions drawn from this study are that captive breeding of crowned lemurs may be enhanced by keeping permanent multi-male, multi-female groups.     

Spatiotemporal distribution of individuals as an indicator for the social system of Lepilemur sahamalaza

Primate social systems are highly diverse, complicating the classification of particularly elusive species that are difficult to observe. The spatial distribution of individuals over time is a critica lindicator for the social organization and long‐term studies are important to establish patterns of social interactions. In recent years, species of the cryptic, nocturnal sportive lemurs of the genus Lepilemur were found to live in pairs in which a single male and a single female share and defend a mutual home range. The present study aimed to forward research into this underrepresented genus by determining the social organization and structure of the Sahamalaza sportive lemur, L. sahamalaza. We collected 773.15 hr of behavioral and GPS data during a period of 10 months (between 2015 and 2016) on 14 individuals: eight females and six males. There was no evidence of pair‐specific home range use as intra‐ and intersexual home range overlap was high. No pattern of social interactions between focal individuals could be distinguished despite high range overlap. Individuals met and interacted infrequently, resulting in an interaction rate of 0.32 interactions/hr. Sleeping associations between adult individuals were never observed. While both sexes had access to multiple potential mating partners, range sizes or ranging distances did not increase in mating periods. Overall, the social system of Sahamalaza sportive lemurs exhibits aspects of a solitary social organization and structure with potential for the polygamous mating system. These findings underline the importance of detailed social ecology studies that can provide the basis for understanding potential environmental influences on social system variability of closely related species.     

The effects of sexual selection and life history on the genetic structure of redfronted lemur, Eulemur fulvus rufus, groups

We examined genetic consequences of basic predictions of life history and sexual selection theory in a wild population of redfronted lemurs. Because group living in lemurs evolved independently from other primates, and because polygynous lemurs deviate in several sexually selected traits from theoretical predictions, data on genetic correlates of their social and mating systems can make important contributions to studies of convergence in social evolution, but such data are not available from wild populations. We extracted DNA from tissue samples obtained from 59 animals living in Kirindy forest, Madagascar, and examined individual variability at several microsatellite loci and the mitochondrial D-loop. We found that closely related females of a single matriline formed the core of the four main study groups. Virtually all haplotypes of adult males differed from those of coresident females, and many male haplotypes were represented by only one or two individuals. Paternity analyses for infants from groups with detailed behavioural data revealed that a disproportionate share of infants were sired by the central, dominant male of a group, despite promiscuous mating. Extragroup paternities were not detected. The skew in male reproductive success cannot be reconciled with the lack of sexual dimorphism and the even adult sex ratios. We therefore conclude that these group-living lemurs converge with many other primates in sex-specific life history trajectories, including female philopatry and male dispersal, but that the observed skew in male reproductive success makes the apparent lack of adaptation to intrasexual selection in certain behavioural, demographic and morphological traits even more puzzling. Copyright 2002 The Association for the Study of Animal Behaviour. Published by Elsevier Science Ltd. All rights reserved.     

A preliminary study of spatial distribution and mating system of pygmy mouse lemurs (Microcebus cf. myoxinus)

According to current hypotheses on the evolution of life history traits and social systems of Malagasy lemurs, nocturnality is associated with a solitary lifestyle and a polygynous or promiscuous mating system. Recent studies, however, have indicated that this may not be true of all lemurs. The goal of this study was to investigate the sociality and the mating system of pygmy mouse lemurs (Microcebus cf myoxinus), which are the smallest known primates, and which retain characteristics of the most primitive primates. I compared my findings with data on the sympatric Microcebus murinus and Cheirogaleus medius. Observational, morphometric and spatial distribution data were obtained by a radiotracking study in 1994, and from a capture-recapture study conducted during 1995/96. Pygmy mouse lemurs usually slept alone in a tangle of vegetation. During the mating season, sleeping sites of males were distributed over a much broader area than were female sites, indicating that male home ranges are larger than those of females. The home ranges of males overlapped during the mating season, and males occasionally roamed over long distances during a single night. Pygmy mouse lemurs forage primarily alone. Analysis of estrus stages indicate that female cycles are unsynchronized during the mating season. There was a lack of sexual dimorphism in body size but not in body mass. Males were heavier than females during the reproductive season but lighter than females the rest of the year. Testes of males varied in size seasonally and enlarged significantly during the mating season. The presence of a vaginal sperm plug in a female indicated the importance of preventing additional matings in this species. Thus pygmy mouse lemurs follow the predictions derived from sexual selection theory for multi-male mating systems with promiscuous matings and male sperm competition.     

Female mate preference to maximize paternal care. II. Female competition leads to monogamy

ABSTRACT A two-step game model of female mate preference and paternal care is examined, with a particular focus on the case of two females and two males. In a mating season, females choose their mates, and in the following breeding season males invest in paternal care, knowing the likelihood of their paternity in chicks. If parental ability is the same between individuals of each sex, the evolutionarily stable mating pattern is always monogamy. If females differ in fecundity and males differ in paternal care capacity, monogamy with assortative mating is likely to be evolutionarily stable. If the male cost function increases at a strongly accelerating rate, however, polyandry is evolutionarily stable when the difference of female fecundity is very large, but the game may have no evolutionarily stable state when the difference of female fecundity is small. The care graph (in which females are connected to males giving paternal care to their chicks) is often much simpler than the mating graph (in which females are connected to males they accepted). To be exact, no "loop" should be included in the evolutionarily stable care graph for the general case of n females and m males. This prediction is in accord with the observed prevalence of social monogamy in spite of genetic promiscuity among altricial birds.     

Paternity and kinship patterns in polyandrous moustached tamarins (Saguinus mystax)

We studied patterns of genetic relatedness and paternity in moustached tamarins, small Neotropical primates living in groups of 1–4 adult males and 1–4 adult females. Generally only one female per group breeds, mating with more than one male. Twin birth are the norm. In order to examine the genetic consequences of this mating pattern, DNA was extracted from fecal samples collected from two principal and six neighboring groups. DNA was characterized at twelve microsatellite loci (average: seven alleles/locus). We addressed the following questions: Do all adult males have mating access to the reproductive female of the group? How is paternity distributed across males in a group? Can polyandrous mating lead to multiple paternity? Are nonparental animals more closely related to the breeders than to the population mean? And, are mating partners unrelated? Breeding females mated with all nonrelated males. In at least one group the father of the older offspring did not sire the youngest infant although he was still resident in the group. We also found evidence for multiple paternity in a supposed twin pair. Yet, within each group the majority (67–100%) of infants had the same father, suggesting reproductive skew. Relatedness within groups was generally high (average R = 0.31), although both nonrelated males and females occurred, i.e., immigrations of both sexes are possible. Mating partners were never found to be related, hence inbreeding seems to be uncommon. The results suggest that while the social mating system is polyandry, paternity is often monopolized by a single male per group. Am J Phys Anthropol, 2005. © 2004 Wiley-Liss, Inc.     

The effects of habitat fragmentation on the social kin structure and mating system of the agile antechinus, Antechinus agilis

Habitat fragmentation is one of the major contributors to the loss of biodiversity worldwide. However, relatively little is known about its more immediate impacts on within-patch population processes such as social structure and mating systems, whose alteration may play an important role in extinction risk. We investigated the impacts of habitat fragmentation due to the establishment of an exotic softwood plantation on the social kin structure and breeding system of the Australian marsupial carnivore, Antechinus agilis. Restricted dispersal by males in fragmented habitat resulted in elevated relatedness among potential mates in populations in fragments, potentially increasing the risk of inbreeding. Antechinus agilis nests communally in tree hollows; these nests are important points for social contact between males and females in the mating season. In response to elevated relatedness among potential mates in fragmented habitat, A. agilis significantly avoided sharing nests with opposite-sex relatives in large fragment sites (but not in small ones, possibly due to limited nest locations and small population sizes). Because opposite-sex individuals shared nests randomly with respect to relatedness in unfragmented habitat, we interpreted the phenomenon in fragmented habitat as a precursor to inbreeding avoidance via mate choice. Despite evidence that female A. agilis at high inbreeding risk selected relatively unrelated mates, there was no overall increased avoidance of related mates by females in fragmented habitats compared to unfragmented habitats. Simulations indicated that only dispersal, and not nonrandom mating, contributed to inbreeding avoidance in either habitat context. However, habitat fragmentation did influence the mating system in that the degree of multiple paternity was reduced due to the reduction in population sizes and population connectivity. This, in turn, reduced the number of males available to females in the breeding season. This suggests that in addition to the obvious impacts of reduced recruitment, patch recolonization and increased genetic drift, the isolation of populations in habitat patches may cause changes in breeding behaviour that contribute to the negative impacts of habitat fragmentation.     

Effects of reproductive and social variables on fecal glucocorticoid levels in a sample of adult male ring-tailed lemurs (Lemur catta) at the Beza Mahafaly Reserve, Madagascar

Glucocorticoids, a group of adrenal hormones, are secreted in response to stress. In male primates, variables such as breeding seasonality, dominance hierarchy stability, and aggressive and affiliative interactions can affect glucocorticoid levels. In this study, we examined interindividual differences in mean fecal glucocorticoid (fGC) levels among males in three groups of wild ring-tailed lemurs to better understand the physiological costs of group living for males in a female-dominant species that exhibits strict reproductive seasonality. Fecal and behavioral data samples were collected during one mating and two postmating seasons (2001 and 2003). The mean fGC levels were examined in relation to reproductive season, male rank, number of resident males, intermale and female-male agonism, and affiliative behavior with females. The mean fGC levels were not significantly elevated during mating season compared to the postmating period. During the mating season, male dominance hierarchies broke down and rank effects could not be tested; however, there was no relationship between male rank and fGC levels in the postmating periods. In 2001, males that resided in the group with the fewest males exhibited lower fGC levels during the postmating period. They also affiliated more with females than did males in the other groups. During the mating season of 2003, males engaged in more affiliative behaviors with females compared to the postmating season, but female-male agonism did not differ by season. However, rates of intermale agonism were significantly higher during mating compared to postmating periods, but such heightened agonism did not translate to a higher stress response. Thus, neither male-male competition for mates nor heightened agonism between males during the breeding season affected male fGC levels. Fewer males residing in a group, however, did have some effect on male-female affiliation and male fGC levels outside of the mating period. Males that live in a group with only a few (two or three) males may experience less physiological stress than those that live in groups with more males.     

Reproductive success in relation to dominance rank in the absence of prime-age males in Barbary macaques

In some primate species dominance rank of males is correlated with reproductive success, whereas in other species this relationship is inconsistent. Barbary macaques (Macaca sylvanus) live in a promiscuous mating system in which males are ranked in a dominance hierarchy that influences their access to females. High-ranking males usually monopolize fertile females during their estrous period and show increased mating activities. Subadult males generally rank below adult males. For Barbary macaque females in the Gibraltar colony, there was no correlation between dominance status and reproductive success. Paternity data for 31 offspring collected over four consecutive breeding seasons were used to test whether male social rank was associated with reproductive success and whether reproductive success was mainly confined to a small number of males. Genetic variation was assessed using 14 microsatellite markers for a dataset of 127 individuals sampled in all five social groups of the Gibraltar colony. Paternity analysis was conducted for offspring in one social group only, where all in-group males were sampled. Eighty-three percent of the offspring could be assigned to an in-group candidate father; none of the extra-group males appeared to have sired an infant. Male dominance rank was not found to contribute to the observed variation in male reproductive output. Fifty-nine percent of the offspring was sired by two low-ranking males, whereas the two top-ranking males sired one-fifth. A highly significant correlation was found for male age and dominance rank. Reproductive success of subadult males might be explained by the gap in the age distribution of male group members. These missing prime males are usually regarded as serious competitors for older males. Subadult males may have gained easier access to females in their absence. In addition, the presence of inbreeding avoidance mechanisms, which might also have overpowered possible rank effects, cannot be excluded.     

Dispersal,Mating Events and Fine-Scale Genetic Structure in the Lesser Flat-Headed Bats

Population genetic structure has important consequences in evolutionary processes and conservation genetics in animals. Fine-scale population genetic structure depends on the pattern of landscape, the permanent movement of individuals, and the dispersal of their genes during temporary mating events. The lesser flat-headed bat (Tylonycteris pachypus) is a nonmigratory Asian bat species that roosts in small groups within the internodes of bamboo stems and the habitats are fragmented. Our previous parentage analyses revealed considerable extra-group mating in this species. To assess the spatial limits and sex-biased nature of gene flow in the same population, we used 20 microsatellite loci and mtDNA sequencing of the ND2 gene to quantify genetic structure among 54 groups of adult flat-headed bats, at nine localities in South China. AMOVA and F _ST estimates revealed significant genetic differentiation among localities. Alternatively, the pairwise F _ST values among roosting groups appeared to be related to the incidence of associated extra-group breeding, suggesting the impact of mating events on fine-scale genetic structure. Global spatial autocorrelation analyses showed positive genetic correlation for up to 3 km, indicating the role of fragmented habitat and the specialized social organization as a barrier in the movement of individuals among bamboo forests. The male-biased dispersal pattern resulted in weaker spatial genetic structure between localities among males than among females, and fine-scale analyses supported that relatedness levels within internodes were higher among females than among males. Finally, only females were more related to their same sex roost mates than to individuals from neighbouring roosts, suggestive of natal philopatry in females.     

Patterns and dynamics of sex-biased dispersal in a nocturnal primate, the grey mouse lemur, Microcebus murinus

We examined predictions on the proportion of dispersing natal males and females, dispersal distances, the age at dispersal and the potential for inbreeding over a 6-year period in a free-living population of grey mouse lemurs. We used monthly mark-recapture procedures to determine individual locations and interindividual distances. The analysis of seven polymorphic microsatellite markers for 213 (130 males, 83 females) individuals allowed us to estimate relatedness coefficients and kinship relationships. Closely related males ranged further from each other than closely related females and natal males were found further from their potential mothers than were females. Natal males were more likely to disperse from their birth sites than females, although male dispersal was not universal. Male breeding dispersal was detected in half of the long-term observations. Males therefore seem to be the predominant vectors for gene flow between populations and social units. Females usually stayed within one to two home range diameters of their potential mother, facilitating the evolution of cooperative behaviour by kin selection among females. Most dispersal took place before the mating season, indicating an age of less than 7 months for natal dispersal. The analysis of spatiotemporal coexistence revealed the potential for inbreeding in only 3.8% of the potential mother-son dyads, but in 21.9% of the potential father-daughter dyads and in 41.7% of other closely related male-female dyads. Copyright 2003 Published by Elsevier Science Ltd on behalf of The Association for the Study of Animal Behaviour      

Social System of <Emphasis Type="Italic">Microcebus berthae</Emphasis>, the World’s Smallest Primate

Our goal was to provide a first characterization of the social system of pygmy mouse lemurs (Microcebus berthae), the worlds smallest primate species. During a 4-mo field study of 12 females and 27 males, we combined capture-recapture and morphometric data with detailed behavioral observations of individually marked subjects and genetic paternity analyses of a population in Kirindy Forest, western Madagascar. We describe the social organization of Microcebus berthae as a solitary forager living in an individualized neighborhood system characterized by extensive intra- and intersexual home range overlap of adult individuals within a male-biased population. Male and female pygmy mouse lemurs inhabited home ranges (males: 4.9 ha; females: 2.5 ha) that are more than twice as large as those of sympatric Microcebus murinus. On average, pygmy mouse lemurs of both sexes spent about half of the days sleeping alone. Preliminary analysis of genetic population structure suggests female philopatry and male dispersal. Sleeping associations of variable composition that consisted not preferentially of close relatives and proximity during part of the nightly activity contributed together with regular social interactions to the maintenance of a social network. The spatial distribution pattern of adult males and females, the absence of sexual size dimorphism, relatively large male testicular volume and moderate female estrous synchrony suggest a promiscuous mating system with a high potential for scramble competition. In general, there are many similarities between the social system of Microcebus berthae and those of other Microcebus spp. However, striking differences with sympatric gray mouse lemurs (Microcebus murinus) in female home range size, dispersion and sleeping behavior indicate the existence of species-specific socioecological adaptations in closely related species occupying very similar ecological niches.     

Courtship Disruptions and Male Mating Strategies: Examples from Female-Defense Mating Systems

Males frequently interrupt the copulation attempts of other males, and these courtship disruptions may limit the extent to which a few males are able to monopolize mating access to females. Males actively defend sexually receptive females in many species in which females form dense aggregations during the breeding season. Across and within such species there is considerable variation in the mating tactics adopted by males, with males in some cases defending groups of females and in other cases sequentially consorting with individual females. Colonial blackbirds have been central to studying this mating system, and we develop a conceptual model for how courtship disruption may account for variation in male mating tactics in this group. Our model assumes that the frequency of disruptions increases with greater colony size. As a consequence, successful copulations are less likely to occur at large colonies than at small colonies, and males are expected to switch from defending multiple females at the colony to consorting individual females away from it. Results from two species of blackbird support the basic assumptions of this model. In one species, the Montezuma oropendola, disruptions occur rarely and males defend groups of females, whereas in the other species, the yellow-rumped cacique, disruptions are frequent and males defend single females. Moreover, consistent with a key prediction, within each species, males associated with small colonies remain at the colony and defend groups of females, whereas males spend little time defending groups of females at large colonies and rarely attempt copulations there. This model has the potential to explain variation in male mating strategies and female monopolization for other taxa in which females form breeding aggregations.     

High levels of genetic monogamy in the group-living Australian lizard Egernia stokesii

The Australian lizard Egernia stokesii lives in spatially and temporally stable groups of up to 17 individuals. We have recently shown that these groups are comprised of breeding partners, their offspring and, in some cases, highly related adults, providing the first genetic evidence of a family structure in any lizard species. Here we investigated the mating system of E. stokesii using data from up to eight polymorphic microsatellite loci and tested the hypothesis that breeding partners are monogamous both within and between mating seasons. Among 16 laboratory-born litters from field collected gravid females from two sites in South Australia, 75% had a single male parent and no male contributed to more than one litter, indicating a high level of genetic monogamy within a season. Additional analyses of field caught individuals, captured between 1994 and 1998, enabled assignment of parentage for 70 juveniles and subadults. These data showed that most young (88.6%) had both parents from within the same group and that high proportions of males (88.9%) and females (63.6%) have multiple cohorts of offspring only with the same partner. Our results suggest that monogamy both within and between seasons is a common mating strategy of E. stokesii and that breeding partners maintain stable associations together and with multiple cohorts of their offspring over periods of up to at least 5 years.     

Intensive genetic assessment of the mating system and reproductive success in a semi-closed population of the mottled sculpin,Cottus bairdi

Most genetic surveys of parentage in nature sample only a small fraction of the breeding population. Here we apply microsatellite markers to deduce the genetic mating system and assess the reproductive success of females and males in an extensively collected, semi-closed stream population of the mottled sculpin fish, Cottus bairdi. In this species, males guard nest rocks where females deposit the eggs for fertilization. The potential exists for both males and females to mate with multiple partners and for males to provide parental care to genetically unrelated offspring. Four hundred and fifty-five adults and subadults, as well as 1,259 offspring from 23 nests, were genotyped at five polymorphic microsatellite loci. Multilocus maternal genotypes, deduced via genetic analyses of embryos, were reconstructed for more than 90% of the analysed nests, thus allowing both male and female reproductive success to be estimated accurately. There was no genetic evidence for cuckoldry, but one nest probably represents a takeover event. Successful males spawned with a mean of 2.8 partners, whereas each female apparently deposited her entire clutch of eggs in a single nest (mean fecundity = 66 eggs/female). On average, genetically deduced sires and dams were captured 1.6 and 9.3 metres from their respective nests, indicating little movement by breeders during the spawning season. Based on a 'genetic mark-recapture' estimate, the total number of potentially breeding adults (c. 570) was an order-of-magnitude larger than genetically based estimates of the effective number of breeders (c. 54). In addition, significantly fewer eggs per female were deposited in single than in multidam nests. Not only were perceived high-quality males spawning with multiple partners, but they were receiving more eggs from each female.     

Androgen levels and female social dominance in Lemur catta

Morphological and behavioural traits which improve agonistic power are subject to intrasexual selection and, at the proximate level, are influenced by circulating androgens. Because intrasexual selection in mammals is more intense among males, they typically dominate females. Female social dominance is therefore unexpected and, indeed, rare. Ring-tailed lemurs (Lemur catta) are sexually monomorphic primates in which all adult females dominate all males. The goal of our study was to test the prediction that female dominance in this species is associated with high androgen levels. Using two captive groups, we collected data on agonistic behaviour and non-invasively assessed their androgen concentrations in faeces and saliva by enzyme immunoassay. We found that adult female L. catta do not have higher androgen levels than males. However, during the mating season there was a twofold increase in both the androgen levels and conflict rates among females. This seasonal increase in their androgen levels was probably not due to a general increase in ovarian hormone production because those females showing the strongest signs of follicular development tended to have low androgen concentrations. At the individual level neither the individual aggression rates nor the proportion of same-sexed individuals dominated were correlated with their androgen levels. We conclude that female dominance in ring-tailed lemurs is neither based on physical superiority nor on high androgen levels and that it is equally important to study male subordination and prenatal brain priming effects for a complete understanding of this phenomenon.